Breeding for provitamin A biofortification of maize (Zea mays L.)

Vitamin A deficiency is widely prevailing in children and women of developing countries. Deficiency of vitamin A causes night blindness, growth retardation, xerophthalmia and increases the susceptibility against epidemic diseases. Among different interventions of overcoming malnutrition, biofortification is the most acceptable and preferred intervention among researchers, growers and consumers. Maize is grown and consumed in those regions where vitamin A deficiency is most prevalent; thus, targeting this crop for provitamin A biofortification is the most appropriate solution. Different breeding strategies including diversity analysis, introduction and stability analysis of exotic germplasm, hybridization, heterosis breeding, mutagenesis and marker‐assisted selection are practised for exploring maize germplasm and development of provitamin A‐enriched cultivars. Genome‐wide association selection and development of transgenic maize genotypes are also being practised, whereas RNA interference and genome editing tools could also be used as potential strategies for provitamin A biofortification of maize genotypes. The use of these breeding strategies for provitamin A biofortification of maize is comprehensively reviewed to provide a working outline for maize breeders.     

Genomic prediction for yields,processing and nutritional quality traits in cultivated potato (Solanum tuberosum L.)

Current potato breeding approaches are hampered by several factors including costly seed tubers, tetrasomic inheritance and inbreeding depression. Genomic selection (GS) demonstrated interesting results regardless of the ploidy level, and can be harnessed to circumvent these problems. In this work, three GS models were evaluated using 50,107 informative SilicoDArT markers and 11 traits in two values for cultivation and use (VCU) potato trials. Two key breeding problems modelled included predicting the performance of (i) new and unphenotyped clones (cross‐validation) and (ii) a VCU using another as training set (TS). GS models performed comparably. Cross‐validation accuracy was high for D35, D45, DMW and BVAL, in ascending order. Prediction accuracies of the VCUs were highly correlated, but the best prediction was obtained for the smaller VCU using the bigger as TS. Cross‐validation and VCU prediction accuracies were higher when bigger TSs were used. The findings herein indicate that GS can be attractively integrated in potato breeding, particularly in early clonal generations to predict and select for traits with low heritability which would otherwise require more testing years, environments and resources.     

Evaluation of the contribution of teosinte to the improvement of agronomic,grain quality and yield traits in maize (Zea mays)

Teosinte, an ancestor to modern maize, displays an excellent performance regarding resistance to stress, but its yield potential has rarely been reported. To evaluate the potential contribution of teosinte to maize improvement, two maize–teosinte backcrossed recombination inbred line (RIL) populations and their corresponding test-cross hybrids were planted for trait assessment. In RN and ZP RIL populations, the average coefficients of variation of 31 agronomic traits were 9.14% (Range, 0.38%‒25.21%) and 6.85% (Range, 0.55%‒27.73%), respectively. The correlation coefficients of 13 common shared traits between RIL populations and test-cross hybrid populations ranged from 0.10 to 0.60 and from 0.06 to 0.72, respectively. A total of 39 and 3 recombined inbred lines, and 29 and 47 test-cross hybrids exhibited higher yields than their checks (RP125, Zheng58, CD189 and ZD958) with the BLUP data, respectively. Furthermore, four test-cross hybrids including RN034/SCML1950, ZP068/Chang7-2, ZP079/Chang7-2 and ZP122/Chang7-2 showed a more stable yield performance, with yield gains of +7.07%, +3.64%, +5.83% and +3.82% over checks, respectively. In conclusion, teosinte could serve as an alien germplasm for maize breeding.     

QTL mapping of maize (Zea mays) stay-green traits and their relationship to yield

A maize genetic linkage map derived from 115 simple sequence repeat (SSR) markers was constructed from an F₂ population. The F₂ was generated from a cross between a stay-green inbred line (Q319) and a normal inbred line (Mo17). The map resolved 10 linkage groups and spanned 1431.0 cM in length with an average genetic distance of 12.44 cM between two neighbouring loci. A total of 14 quantitative trait loci (QTL) were detected for stay-green traits at different postflowering time intervals and identified by composite interval mapping. The respective QTL contribution to phenotypic variance ranged from 5.40% to 11.49%, with trait synergistic action from Q319. Moreover, maize stay-green traits were closely correlated to grain yield. Additional QTL analyses indicated that multiple intervals of stay-green QTL overlapped with yield QTL.     

New techniques for breeding maize (Zea mays) varieties with fall armyworm resistance and market-preferred traits for sub-Saharan Africa

Deploying maize varieties with fall armyworm (Spodoptera frugiperda [J.E. Smith]; FAW) resistance, desirable product profiles (PPs) and climate resilience is fundamental for food and economic security in sub-Saharan Africa (SSA). This study reviewed and identified challenges and opportunities for effective and accelerated breeding of demand-led maize hybrids with FAW resistance and adaptation to the diverse agro-ecologies of SSA. Lessons were drawn on improving breeding efficiency through adequate genetic variation delivered via prebreeding programmes, speed breeding and a reduced breeding stage plan. Appropriate PPs aligned with demand-led breeding approaches were highlighted as foundations for variety design and commercialization. Challenges to accelerated FAW resistance breeding in maize included inadequate funds and modern tools; poor adaptation of some exotic donor parental lines; lack of information on FAW resistance among local varieties; lack of integration of molecular markers associated with FAW resistance and agronomic traits into selection plans; and limited infrastructure for FAW rearing and germplasm screening. Integration of modern breeding tools and scientific innovations were recommended for accelerated development and release of FAW resistant and market-preferred maize varieties.     

A QTL atlas for grain yield and its component traits in maize (Zea mays)

Grain yield and its component traits are essential targets in maize breeding. These traits are genetically complex and controlled by a large number of quantitative trait loci (QTL). The aim of this study was to compile reported QTL and major genes for grain yield and its component traits in a QTL atlas, as a valuable resource for the maize community. To this end, 1,177 QTL related to maize yield were collected from 56 studies published between 1992 and 2018. These QTL were projected to genetic map “IBM2 2008 Neighbors”, which led to the identification of 135 meta-QTL. Some genomic regions appear to be hotspots for yield-related meta-QTL, often affecting more than one of the investigated traits. Moreover, we catalogued 20 major maize loci associated with yield and identified 65 maize homologs of 21 rice yield-related genes. Interestingly, we found that a significant proportion of them are located in meta-QTL regions. Collectively, this study provides a reference for QTL fine-mapping and gene cloning, as well as for molecular marker-assisted breeding of yield-related traits in maize.     

Farmers' highland maize (Zea mays L.) selection criteria: Implication for maize breeding for the Hararghe highlands of eastern Ethiopia

Despite the presence of large numbers of improved maize cultivars, farmers in the Hararghe highlands of eastern Ethiopia persistently grow local cultivars and are not benefiting from the varietal improvement program. By growing local cultivars farmers obtain an average yield of 1.2 t ^-1 whereas research has released cultivars yielding 5–11 t ^-1under on-station conditions. Recognizing this and the important role maize is playing in the livelihood system of farmers in eastern Ethiopia; Participatory Varietal Selection (PVS) was conducted for three consecutive seasons (1996–1998) in three locations. The objectives of the study were to identify farmers' cultivar selection criteria for future breeding, to enable farmers to assess the performance of improved cultivars under their management, to increase farmers' access to the cultivars of their preference by injecting source seed into the local seed system, and to investigate whether breeding for wide adaptation like for the Hararghe highlands has any drawbacks or not. The study indicated farmers' maize varietal selection criteria together with the differences in selection criteria across locations classified under the same adaptation zone. Though farmers selected some of the many improved cultivars tested, no improved cultivar had all the characteristics, which farmers want in a single cultivar. To be able to combine cultivar selection traits farmers considered cultivar combinations in all locations, but not the single `best' cultivar. Despite the yield advantage of hybrids, farmers selected Open Pollinated Varieties (OPVs) because they had more preferred traits. Farmers also preferred to retain their local cultivars despite their lower yield compared to most of the improved cultivars. Both situations confirmed how resource poor farmers' cultivar requirements are much more diverse than yield per se. Gender and social group-driven difference in cultivar preference was also observed. The study revealed the need for proper zoning of the Hararghe highlands as a prerequisite to developing better adapted maize cultivars to the varied agro ecological and socioeconomic niches. As shown in this study, proper zoning, due consideration to farmers' relevant selection traits and wider use of participatory approaches should be adapted to develop cultivars which can gain farmers acceptance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genomic prediction of the general combining ability of maize lines (Zea mays L.) and the performance of their single crosses

The objective of this study was to assess the effectiveness of genomic selection (GS) on predicting the general combining ability (GCA) of maize lines and the performance of their single crosses. Eight maize lines developed from the different self‐pollination generations of Chalqueño race, along with their 24 single crosses, were evaluated in the field during the years of 2011, 2012 and 2013. Genomic prediction results using genotyping‐by‐sequencing‐based single nucleotide polymorphisms showed that the GCA classification of the parental lines estimated from the SNP information was consistent with the phenotypic classification of the lines evaluated from the field trial data. The prediction accuracy values estimated from the cross‐validation method ranged from 0.49 to 0.61 in the different prediction models. Yield performance of the unevaluated single crosses was predicted based on their SNP information. The total genetic variance of the yield of the single crosses was most explained by the GCA effects. Compared with phenotyping method, GS is a more effective and efficient approach to predict the GCA of maize lines and their hybrid performance.     

Genetic analyses of agronomic traits in Tartary buckwheat (Fagopyrum tataricum (L.) Gaertn.)

The consumption of products made from Tartary buckwheat (Fagopyrum tataricum (L.) Gaertn.) has increased in recent years in Japan. Increased consumer demand has led to recognition of the need for early varieties of this crop with high and stable yields. In order to accomplish this, more information is needed on the genetic mechanisms affecting earliness and yield. We conducted genetic analysis of 3 agronomic traits (days to flowering, plant height and total seed weight per plant) to segregate F₂ and F₃ populations derived from a cross between Tartary buckwheat cultivars ‘Hokuriku No. 4’ and ‘Ishisoba’. Broad-sense heritability estimates for days to flowering, plant height and total seed weight were 0.70, 0.62 and 0.75, respectively, in F₃ population. Narrow-sense heritability for total seed weight (0.51) was highest, followed by heritability for days to flowering (0.37), with heritability for plant height (0.26) lowest. Later flowering was associated with increased plant height and higher yields. From the F₄ generation, we identified twelve candidate plants with earlier maturity and reduced plant height compared to ‘Hokuriku No. 4’, but almost the same total seed weight. These results suggest that hybridization breeding using the single seed descent (SSD) method is an effective approach for improving agronomic characteristics of Tartary buckwheat.     

Genomic selection in hybrid breeding

While hybrid breeding is widely applied in outbreeding species, for many self‐pollinating crop plants, it has only recently been established. This may have had its reason in the limitations of methods available for hybrid performance prediction, in particular when established heterotic pools were absent. Genomic selection has been suggested as a promising approach to resolve these limitations. In our review, we briefly introduce the principles of genomic selection as an extension of marker‐assisted selection using genome‐wide high‐density molecular marker data and discuss the advantages and limitations of currently used algorithms. Including the outcome from a recent extended approach to hybrid wheat as a timely example, we summarize current progress in empirical studies on the application of genomic selection for prediction of hybrid performance. Here, we put emphasis on the factors affecting the accuracy of prediction, pointing in particular to the relevance of relatedness, genotype x environment interaction and experimental design. Finally, we discuss future research needs and potential applications.     

Breeding for yield, in mixtures of common beans (Phaseolus vulgaris L.) and maize (Zea mays L.)

Summary Despite the growing industrialization, technification and transformation that is happening in the agriculture around the world, and despite that agricultural research has always concentrated its effort on sole crops, multiple cropping systems have historically been important for common bean production in tropical countries. The reasons for this fact, are economical and social, as well as biological. Bean breeders have always been questioned on their work, because the development of new varieties is usually done in sole crop, but the varieties are grown in either systems. This paper addresses a set of questions that are usually presented to the breeders, in light of the evidence obtained from many trials conducted in Brazil and in the U.S.A.: Will the genotypes bred for sole crop conditions, perform well when grown in intercrop; How different should a genotype be, for cultivation in intercropping compared to genotypes developed for sole crop conditions; Is there a need for special breeding programs for intercropping and How could a breeding program focus the question of multiple (associated) cropping?     

Genetic variation of seedling traits responded to brassinosteroid and gibberellin inhibitors in maize (Zea mays) doubled haploid lines

The purpose of this study was to examine the genotypic variation in maize doubled haploid (DH) lines response to brassinosteroid and gibberellin inhibitors. Plant responses to hormone inhibitors were determined in growth chamber experiments using germination paper for three different seedling treatments: application of propiconazole (Pcz), uniconazole (Ucz) or water (control). Mesocotyl length (ML) was more sensitive to hormone inhibitors, especially to the Ucz treatment, than other seedling traits. ML was significantly correlated with other traits in the Ucz treatment. All the seedling traits showed moderate-to-high broad sense heritability values, ranging from 0.39 to 0.82. The Euclidian genetic distances of inbred line pairs ranged from 1.27 to 19.94, indicating there was a high level of variability across the maize DH lines used in this study. DH lines with extreme MLs were identified, which can provide valuable breeding resources for improving abiotic stress tolerance, and for further genetic studies.     

The alternative breeding approaches for improving yield gains and stress response in pigeonpea (Cajanus cajan)

Pigeonpea is an important food legume crop of semi‐arid tropical regions. Plateauing of pigeonpea yield has been worrying breeders for the past 6–7 decades. Serious breeding efforts made during this period resulted in various high‐yielding and disease resistant cultivars. However, the gains in pigeonpea productivity have been modest. The authors, while reviewing this situation, conclude that long generation turnover, complexity of biological traits, low selection response and overreliance upon pedigree breeding present the key bottlenecks for this situation. In this paper, some alternative breeding approaches and technologies are suggested for the genetic enhancement of yield stability and stress response of pigeonpea.     

Effect of pollen storage by drying and deep-freezing on the expression of different agronomic traits in maize (Zea mays L.)

Summary Pollen from the inbred maize line HMv 1645 was used to study the effect of pollen treatments (drying and deep-freezing) on the phenotypic performance of the next generation. Fresh and artificially dried pollen samples with different water contents (56%, 18%, 13% and 10%) were used for sib pollinations immediately after collection or drying. Samples containing low amounts of water were then stored in liquid nitrogen for 7 days. Fertilization ability of the samples with 13% water was the highest after storage. Plant characteristics of the next generation originated from the seeds set by differently treated (fresh, dried to 13% water and deep-frozen) pollen were examined and statistically analysed. Pollen treatments due to the pollen storage procedure did not cause detectable changes in quantitative characters of the next generation.     

Genetic relatedness and the ratio of subpopulation‐common alleles are related in genomic prediction across structured subpopulations in maize

Genomic prediction (GP), which could predict the breeding value of crop plants genotyped with molecular markers, has been carried out in multiple species. Prediction accuracy (PA) of GP depends on various factors, including genetic relatedness and genetic basis. In this study, we examined the rationale for the low PA of GP when the training and validation populations were distinct using 170 temperate inbred lines and 210 tropical and subtropical inbred lines, respectively. All inbred lines were evaluated for 17 traits and genotyped with 550K high‐density markers. The results show that: (a) the influences of heritability and marker number on PA reflected variations in phenotypic variance captured by the genetic information; (b) the low PA of GP when the training and validation populations represent structured subpopulation is related to the ratio of subpopulation‐common alleles (RSCA) and the genetic relatedness between the two subpopulations; (c) RSCA and PA increased with the increase of genetic relatedness, suggesting that these three factors were related. Our findings would provide references when performing GP, and guidance when designing breeding populations.     

Recurrent selection for seed quality traits in an opaque-2 maize population

Summary The aim of this research was to evaluate the response to phenotypic recurrent selection for four seed quality traits, i.e. protein content, dye binding capacity, tryptophan content, and specific weight, in a modified opaque-2 population of maize (Zea mays L.). Changes in other agronomic traits were also studied. Four selection cycles, using as selection criterion an index constructed from the four seed quality traits, were completed on an individual plant basis. The material from each cycle and from testerosses of each cycle with a homozygous opaque-2 tester was grown for two consecutive years to evaluate the progress due to selection. The results showed that four cycles of recurrent selection induced sizeable variations among cycles for all selected traits. The rate of progress for the four seed quality traits was nearly linear. Further-more, theldata show that improvement in seed-protein related traits had no negative effects on grain yield and kernel weight, when evaluated as the mean performance of testerosses. There was also no appreciable alteration in the other agronomic traits caused by selection for seed quality traits.     

Quantitative trait loci mapping of maize (Zea mays) ear traits under low-phosphorus stress

We constructed an F_2:3 population of 180 individuals from a cross between the maize genotypes 082 (high phosphorus use efficiency) and Ye107 (low phosphorus use efficiency). We used the population to perform quantitative trait loci (QTL) mapping for eight ear traits (ear diameter [ED], ear length [EL], ear weight [EW], ear kernel weight [EKW], row number per ear [ERN], bald tip length [BTL], kernel number per row [RKN] and number of seeds per plant [NSP]) in three low-phosphorus environments. A total of 36 QTL for ear traits were detected in at least one of three single environments and were located on all maize chromosomes except for chromosome 10; the explained phenotypic variation ranged from 5.91% to 12.80%. The QTL JAqEKW5-1, JAqEL3-1, JAqEW1-1, JAqBTL1-1 and JAqNSP5-2 were identified and stably expressed in single-environment and joint-environment analyses. Some novel QTL that have not been reported previously were also detected. Most of the identified QTL had stable effects in all low-phosphorus environments, suggesting that they may be useful for molecular breeding to develop low-phosphorus tolerant maize varieties.     

植物基因组选择及其应用研究进展

为促进基因组选择在植物遗传改良中的应用,笔者总结了基因组选择的原理与方法、分析了基因组选择育种的统计方法,指出了各种因素对基因组选择的影响,总结了基因组选择面临的挑战,并讨论了基因组选择在植物数量性状分子育种研究中可能的应用。认为随着基因型分析成本的降低和统计方法的发展,植物基因组选择将会逐步完善,将在植物基因组育种中发挥重要的作用。